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aqua, International Journal of Ichthyology

Fishes of Bermuda

William F. Smith-Vaniz1 and Bruce B. Collette2

1) Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611 USA. Email: [email protected] 2) National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Smithsonian Institution, Washington D.C. 20560 USA. Email: [email protected]

Received: 13 June 2013 – Accepted: 21 September 2013

Abstract Résumé Here we add a recently described species of Hyporham- Ici, nous ajoutons une espèce récemment décrite d’Hy- phus (Hemiramphidae), recognize Chromis bermudae porhamphus (Hemirhamphidae), reconnaissons la validité (Pomacentridae) as valid, and remove Parasphyraenops atri- de Chromis bermudae (Pomacentridae) et ôtons Para- manus (Serranidae) from the list of Bermuda endemics, sphyraenops atrimanus (Serranidae) de la liste des endé- changing the total number of Bermuda endemic fishes to miques des Bermudes, en portant le nombre total de pois- seven species, eight if Clepticus sp. (Labridae) proves to be sons endémiques des Bermudes à sept espèces, huit si Clep- new, excluding from consideration several land-locked ticus sp. (Labridae) s’avère une espèce nouvelle, en ne te- species of Fundulus. First Bermuda records are docu- nant pas compte des espèces de Fundulus peuplant les ter- mented for 24 species but five others, Carcharodon car- res. Les premiers relevés des Bermudes signalent 24 charias, Hyporhamphus unifasciatus, Rypticus subbifrenatus espèces, mais cinq autres, Carcharodon carcharias, Clepticus parrae and Eleotris pisonis, based on misidentifi- Hyporhamphus unifasciatus, Rypticus subbifrenatus, Clepti- cations, are removed. Biological or distributional notes are cus parrae et Eleotris pisonis sont écartées suite à des identi- included for 22 species and nomenclatural changes apply fications erronées. Des notes sur la biologie et la distribu- to another 12 species. Reproductions of recently discov- tion sont incluses pour 22 espèces et les modifications de ered historical watercolors or pencil sketches by Colonel nomenclature appliquées à 12 autres espèces. Des repro- H. W. Drummond-Hay are provided for 14 species of ductions d’aquarelles historiques ou d’esquisses au crayon, Bermuda fishes. récemment découvertes, du Colonel H. W. Drummond- Hay sont présentées et concernent 14 espèces de poissons Zusammenfassung des Bermudes. Wir fügen hier eine kürzlich beschriebene Art der Gat- tung Hyporhamphus (Hemiramphidae) hinzu, anerkennen Sommario Chromis bermudae (Pomacentridae) als gültig und entfer- In questo articolo con l'aggiunta di una specie recente- nen Parasphyraenops atrimanus (Serranidae) aus der Liste mente descritta di Hyporhamphus (Hemiramphidae), il der endemischen Arten Bermudas. Damit ändert sich die riconoscimento di Chromis bermudae (Pomacentridae) come Zahl der endemischen Fische Bemudas zu sieben Arten valida e la rimozione di Parasphyraenops atrimanus (Ser- oder acht, falls Clepticus sp. (Labridae) als neu anerkannt ranidae) dalla lista delle specie endemiche delle Bermuda, il werden kann; einige ans Land gebundene Fundulus-Arten numero totale di queste specie, escludendo dal conteggio bleiben dabei außer Betracht. Erstnachweise für die quelle d’acqua dolce di Fundulus, viene portato a sette, even- Bermuda werden von 24 Arten dokumentiert, während tualmente otto se Clepticus sp. (Labridae) si rivelerà come fünf Arten wieder gestrichen werden, da ihre angeblichen una nuova specie. È documentata per la prima volta la pre- Nachweise auf Fehlbestimmung beruhten, nämlich: Car- senza di 24 specie, ma altre cinque, Carcharodon carcharias, charodon carcharias, Hyporhamphus unifasciatus, Rypticus Hyporhamphus unifasciatus, Rypticus subbifrenatus, Clepticus subbifrenatus, Clepticus parrae und Eleotris pisonis. Für 22 parrae e Eleotris pisoni, sulla base di errori di identificazione, Arten werden Anmerkungen über Biologie und Verbrei- vengono rimosse. Per 22 specie sono incluse note biologiche tung angefügt, zu weiteren 12 Arten sind Änderungen der o di distribuzione, mentre cambiamenti della nomenclatura Nomenklatur vermerkt. Außerdem werden kürzlich sono applicate ad altre 12 specie. Infine, sono presentate wiederentdeckte historische Aquarell- oder Bleistiftskizzen riproduzioni di schizzi a matita e acquerelli storici, recente- von Colonel H. W. Drummond-Hay zu 14 Bermuda- mente ritrovati, di 14 specie di pesci delle Bermuda eseguiti Fisch arten wiedergegeben. dal colonnello H. W. Drummond-Hay.

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INTRODUCTION Newly discovered Drummond-Hay paintings: The purposes of this paper are to update our Colonel Henry M. Drummond-Hay’s unpublished Fishes of Bermuda (Smith-Vaniz et al. 1999), to notes and watercolors of Bermuda fishes (1847-51) revalidate Chromis bermudae and to publish addi- have great scientific value as the first attempt to tional illustrations made by Col. Drummond-Hay accurately document the fishes of Bermuda. Our recently discovered in an on-going inventory of book included the first color reproductions of ichthyological illustrations under way at the Smith- Drummond-Hay paintings (Plates 1-9, depicting sonian Institution by Lisa Palmer. 41 species) and we documented (Table II) the exis- Unfortunately, a number of recent authors have tence of 76 of his whole-fish illustrations (exclud- failed to check distributions of some fishes against ing duplicates) of 70 species, housed at either the our book where we add new records, validate many Smithsonian Institution or the Bermuda Govern- previous records, and give reasons for not accepting ment Archives. As we discussed in our original bio- some historical records of fishes from Bermuda. In graphical notes (modified excerpts reproduced a review of the recent FAO guide to western cen- below with permission, courtesy of the American tral Atlantic living marine resources (Carpenter Society of Ichthyologists and Herpetologists), 2003), Gilbert (2005) provided lists of species Drummond-Hay made a duplicate set of his 100 whose distributions differed from those given in watercolors of Bermuda fishes but no list of these our book and the FAO guide. He listed 32 species paintings has been located. Among the missing we verified as occurring in Bermuda that were not watercolors, is one of a beautiful 52.5 lb grouper included on the FAO distribution maps. Another (local name “Guinea chick hamlet”) caught on the 19 species that we documented from Bermuda outer reef in 1851. G. Brown Goode and Tarleton were not included in the FAO accounts. Four H. Bean (both of whom independently visited the species undocumented from Bermuda and not islands and made major contributions to Bermuda shown on the FAO distribution maps were listed as ichthyology) had an opportunity to examine that occurring in Bermuda in the FAO accounts and particular painting and were so impressed by it that three others with unverified Bermuda occurrences they subsequently (Goode and Bean 1878) named were indicated as present there on the FAO distri- the species Epinephelus drummondhayi. Unfortu- bution maps. Five additional species were listed as nately, this grouper is now listed as Critically occurring in Bermuda in the FAO accounts but are Endangered throughout its range (Sadovy de not considered to be present there. Mitcheson et al. 2013). Although this painting has not been located, we here document the recent dis- METHODS AND MATERIALS covery of additional original watercolors (Figs 3- We present comments, corrections, and additions 15) and two pencil sketches (Figs 4 and 16), now following the order used in our book. Addition of accounting for a total of 84 species of Bermuda the Lampridae is given in phylogenetic order fol- fishes. As an example of the scientific value of these lowing Nelson (2006). Scientific names of taxa, watercolors and sketches, see following accounts of authors and dates of original description follow Haemulon macrostomum under Haemulidae and recent on-line versions of the Catalog of Fishes Masturus lanceolatus under Molidae. (Eschmeyer 2013). Specimen lengths are given in Henry Maurice Drummond-Hay (1814-1896): mm as standard length (SL), fork length (FL) or Colonel Drummond-Hay (Figs 1-2), son of Sir total length (TL). Institution collection abbrevia- Adam Drummond, K. C. H., Admiral, R. N., and tions are as follows: Academy of Natural Sciences, Lady Charlotte Murray, was born at Bath, Eng- Philadelphia (ANSP); Bermuda Aquarium, Nat- land; family seat at Megginch Castle, Perthshire, ural History Museum and Zoo (BAMZ); Museum Scotland. When he married Charlotte Elizabeth, of Comparative Zoology, Harvard University only daughter of James Hay of Segginch, in Octo- (MCZ); University of Michigan Museum of Zool- ber 1859, he changed his name to Drummond- ogy, Ann Arbor (UMMZ); National Museum of Hay. He was educated by private tutor at Meg- Natural History, Washington, D.C. (USNM); ginch and then boarded at Finchley Manor outside Humboldt-Universität, Museum für Naturkunde, London but received no university instruction. In Zoologisches Museum, Ichthyologie, Berlin 1832 he also studied taxidermy for six months (ZMB); and Zoological Museum, University of under Henry Linder, curator of the Museum at Copenhagen (ZMUC). Geneva, and a leading authority on the ornithol-

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ogy of Switzerland and the Alps. He joined the species accounts that pertain to some of the 42nd Royal Highlanders (the Black Watch) after Bermuda Archives’ watercolor holdings are avail- the successful purchase of an ensigncy by his father able, but the status of most of the Drummond-Hay in 1832. During his 20-year military career, Drum- ichthyological manuscript is unknown. Among his mond-Hay was promoted to Lieutenant in 1837 little known accomplishments is apparently having and to Captain in 1842; he retired from active ser- first pioneered the sport of fishing for needlefish vice in June 1852 and became a Lieutenant with a fly rod. In his unpublished hand-written Colonel (1855-1870) of the volunteer Royal notes on Tylosurus acus, apparently intended to Perthshire Rifles, Scotland; he commanded the accompany his beautiful watercolor of the head of regiment during the Crimean war, retiring as a full this large needlefish (our original Plate 2-27), he colonel in 1872. His army postings included: wrote: Corfu in 1835; Glasgow and Dumbarton 1837-8; “This large species of Garfish is very aptly named Ireland 1838-9; Corfu 1841-3; Malta 1843-7; the Hound fish by the Bermudians, as they may Bermuda 1847-51; and Halifax, Nova Scotia incessantly be seen like a pack of hounds in chase 1851-2. of large schools of fry, which in their hurry and He was Commodore of the Royal Bermuda Yacht alarm jump out of the water skipping and scudding Club from 1849-1851. In addition to his artistic before their pursuers in a rolling mass over the abilities, he was a keen naturalist and combined his smooth surface. Taking advantage of the Hound- enthusiasm for fishing and hunting with describing fishes predatory habits, I used often to fish for natural history. Drummond-Hay reportedly con- them in the cut between Harrington Sound and tinued his drawings and notes on Bermuda fishes the flats with a good strong salmon rod and tackle in Halifax, his next duty station after Bermuda. ….using a white fly sunk with a no. 1 shot a little These appear to have been made in preparation for below the surface, casting and working the fly publication and were said to have filled two large much the same way as salmon fishing. This they manuscript volumes. Copies of original single page took readily, affording splendid sport, especially if

Fig. 1. Henry Maurice Drummond-Hay (1814-1896), Fig. 2. Henry Maurice Drummond-Hay (1814-1896), about 1838. Courtesy of Perth Museum & Gallery, Perth about 1890. Courtesy of Perth Museum & Gallery, Perth & Kinross Council, Scotland. & Kinross Council, Scotland.

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a large one, which would run and dance out of the Hangover Hole dive site, 32°14’52.54”N, water like a clean run grilse reminding one of fishing 64°48’58.40”W on April 22, 2013. They esti- on some rapid running salmon river in Scotland. I mated that it was about 8-9 ft long from snout to am told that following my example (the first who tail tip with a wingspan of about 5-7 ft. We include tried it) that rodfishing for them has since become one of several photographs (Fig. 17) taken by quite a pastime for the officers of the garrison”. Gunar Mayer. Identification was confirmed by Primarily interested in ornithology, Drummond- John McEachran. This is the first Bermuda record Hay also compiled notes on his observations of the of this stingray which is found from Cape Cod to Mediterranean avifauna, which were published in Argentina and also in the eastern Atlantic the Annals & Magazine of Natural History in 1843. (McEachran & de Carvalho 2003). A subsequent publication appeared following a Elopidae month-long surveying trip to Tunis aboard Elops smithi McBride, Rocha, Ruiz-Carus, and “H.M.S. Beacon” in 1845. On his return to Bowen 2010, Southern Ladyfish. Based on exami- Europe in December 1852, Drummond-Hay nation of a large number of leptocephalus larvae, observed a Great Auk on the edge of the New- Smith (1989) pointed out that there were two foundland banks at close range with his field forms, probably different species, of Elops in the glasses, and he was fond of relating that he believed western Atlantic. A high-count morph (79-86 total he was the last person to have seen a living exam- myomeres) which he identified as Elops saurus and ple of that great bird. Upon retirement from the a low-count morph (74-78 total myomeres) which army, he became the first president of the British he felt probably represented an undescribed Ornithological journal The Ibis. Drummond-Hay species. The single Bermuda specimen that we was also an active member of the Perthshire Soci- examined has 73 vertebrae and so now can be iden- ety of Natural Science for 25 years, serving as its tified as E. smithi (McBride et al. 2010). president for two terms, 1882-1884, and thereafter Ophichthidae as Honorary Curator of the Society’s museum. For Ophichthus ophis (Linnaeus 1758), Spotted Snake the last twenty years of his life his interests were Eel. This eel, which attains at least 192 cm TL, was focused on the natural history of Perthshire, and first documented from Bermuda based on a Drum- his contributions during that time were mostly mond-Hay watercolor of a “Panther eel” (our Pl. 1- published in the Society’s Proceedings or in the 21) collected in September 1849. The following Annuals of the Scottish Naturalist. two records are the first reports of adults of this eel in over 70 years. On September 23, 2009, a beach- Notes on Bermuda fishes comber reported a moribund and unusual spotted Lamnidae snake eel (Fig. 18) from Clearwater Beach in St. Carcharodon carcharias (Linnaeus 1758), White Davids during a prolonged fish kill event (S.R. Shark. We reported the presence of the White Smith, pers. comm.). The 167 cm TL specimen Shark in Bermuda based on identification of a pho- has been catalogued as BAMZ 2009-265-013. tograph (our Fig. 92) but admitted the possibility Lucas (2012) also includes a photograph on this that it might have been a Shortfin Mako. Castro eel, discovered by Russell Whayman near South- (2011:259), who we did not know was on the boat west Breaker, with its head partially protruding when the photograph was taken, identified a shark from the sand. caught later that day a mile or so from where the Bermuda individuals of this eel have unusually photograph was taken as an unusually large Short- small body spots but agree in other characters of the fin Mako; accordingly, there are no verified records species. John E. McCosker confirmed the identifica- of White Shark from Bermuda. tion. This large ophichthid is also known from Isurus oxyrinchus Rafinesque 1810, Shortfin Mako. North Carolina to Florida and isolated collections We reported several records of the Shortfin Mako from the West Indies to southern Brazil, and the from Bermuda and now correct the previous record eastern Atlantic Ocean (McCosker et al. 1989). of a White Shark as another Shortfin Mako record. Engraulidae Dasyatidae Anchoa choerostoma (Goode 1874), Bermuda Dasyatis centroura (Mitchill 1815), Roughtail Anchovy. Neighbor-joining trees based on both the Stingray. A female Roughtail Stingray was found control region and ITS1 sequences show that the by Christopher Brown and Gunar Mayer at the endemic Anchoa choerostoma is more closely related

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to A. mitchilli, whose range is restricted to the 1,600 m but large individuals have been found Atlantic and Gulf coasts of the United States than floating at the surface a number of times (Cohen et it is to A. hepsetus, whose range extends throughout al. 1991). the Caribbean (Johnson 2003). Bythitidae Lampridae Ogilbia cayorum Evermann & Kendall 1898, Key Lampris guttatus (Brünnich 1788), Opah. Opahs Brotula. A recent revision of Ogilbia by Møller et (genus Lampris) are epi-mesopelagic fishes found al. (2005) confirms that there is only one species of in all oceans. Two species are currently recognized the genus in Bermuda. as valid: Lampris immaculatus Gilchrist 1904, an Antennariidae unspotted Indo-West Pacific species, and the spot- Col. Drummond-Hay illustrated two species of ted L. guttatus. The latter species is considered to frogfishes of the genus Antennarius, A. multiocella- have a circumtropical distribution but this is tus (Valenciennes 1837) and A. scaber (Cuvier uncertain because there may be several spotted, 1817), the latter considered by many to be a junior cryptic species (Hawn & Collette 2012). We have synonym of A. striatus (Shaw 1794). Frogfishes examined three color photographs of spotted opahs have a remarkable ability to alter their coloration to from the vicinity of Bermuda, only one with col- match the surrounding background. We incor- lection data. That individual (Fig. 19), which rectly thought that Drummond-Hay had illus- weighed about 55 lbs was caught 29 April 2008 on trated two color phases of A. scaber based on spec- a longline about 30 mi. north of Bermuda by com- imens caught on 28 Oct. and 8 Dec. 1847, respec- mercial fisherman David Soares. tively, which he referred to as the “orange” and Synodontidae “striped toad fish”. However, in his unpublished A recent important paper on lizardfishes of the notes (quoted on p. 160) he had correctly recog- genus Synodus (Frable et al. 2013) describes one nized that the “orange” and “black toad fish” are new species, resurrects another one from syn- two color phases of the species now known as A. onymy, and redescribes several easily misidentified multiocellatus. Thus the legend for our color plate Caribbean species. That publication does not alter 2-23 should have been labeled as A. multiocellatus, the taxonomic concept or nomenclature of any of not A. scaber. the Bermuda species. Antennarius scaber (Cuvier 1817), Splitlure Frog- Carapidae fish. Since the illustrations by Col. Drummond- Carapus bermudensis (Jones 1874), Pearlfish. The Hay were not of this species as noted above, a pho- first specimen collected in Bermuda (Fig. 20) since tograph (Fig. 21) of a recently collected specimen its original description in 1874 came out of a sea (BAMZ 2010-267-017) by Kimberly Holzer is of cucumber, Actinopyga agassizi when it was cut open interest. by Wolfgang Sterrer in February 2002 (Sterrer Antennatus bermudensis (Schultz 1957), Island 2002a). This confirms Dr. Sterrer’s prediction that Frogfish. In a recent molecular revision of the the host of Carapus, at least in Bermuda, is the rare Antennariidae (Arnold & Pietsch 2012), this nocturnal Actinopyga rather than the more com- species was moved into the A. nummifer species mon Isostichopus. Two specimens are preserved at group in the genus Antennatus Schultz 1957. BAMZ, 2002-201-006 and 2002-203-007. In Fowlerichthys ocellatus (Bloch & Schneider 1801), addition, three vexillifer larvae were collected near Ocellated Frogfish. Arnold & Pietsch (2012) Bermuda by the Ocean Acre program and are cat- assigned Antennarius ocellatus to the genus Fow- alogued as follows: USNM 218380 (1), Ocean lerichthys Barbour 1941. Acre 12-34A; USNM 218381 (1), Ocean Acre 12- Mugilidae 6M; USNM 218382 (1), Ocean Acre 12-1A. Mugil curvidens Valenciennes 1836, Dwarf Mullet. Ophidiidae In their appendix on p. 96 (Harrison et al. 2007) Lamprogrammus brunswigi (Brauer 1906). A listed a single (180 mm SL) specimen: MCZ 17548 headless specimen was found floating at the sur- from “Bermuda, north Sargasso Sea, no collector or face. Based on the photograph, Drs. D. M. Cohen date”. This record documents the fourth species of and J. Nielsen agree that it is one of the giant Lam- mullet in Bermuda. The Dwarf Mullet is also known programmus species, probably L. brunswigi. This is from Ascension Island, the Bahamas, Antilles, and a circumtropical species (except absent from the south to Rio de Janeiro, Brazil (Harrison 2003) so its eastern Pacific) usually found at depths of 800- occurrence in Bermuda is not surprising.

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Belonidae ative to keep fish from different ponds separated Tylosurus crocodilus (Peron & Lesueur 1821), and manage the various ponds or threatened pop- Houndfish. Although all previous museum speci- ulations as discrete units. Accordingly, a recovery mens from Bermuda identified as T. crocodilus were plan for Bermuda killifish has been proposed (Out- misidentifications of T. acus acus, we can now add erbridge & Sarkis 2012). T. crocodilus to the fauna of Bermuda based on the During an 18-month study in 2004-2005, collection of two adults. John Galbraith collected researchers undertook surveys using a combination one (USNM 385990) on hook and line at Non- of direct observation and baited trapping to esti- such Island on Aug. 16, 1999. A second specimen mate abundance, size frequencies, and sex ratios (USNM 385891) was also caught on hook and line among the isolated killifish populations (Outer- by James Liao on Aug. 14, 2001 off the rocks along bridge et al. 2006; Outerbridge et al. 2007a). Mark the railroad trail just north of Whalebone Bay. and recapture studies produced the following esti- Hemiramphidae mates of population size: Mangrove Lake (11,325 Hyporhamphus collettei Banford, 2010, Collette’s ± 1,884 fish), Trott’s Pond (7,926 ± 1,576), Lover’s Halfbeak. The Bermuda population of Hyporham- Lake (8,508 ± 1,347), Blue Hole Bird Pond (5,394 phus has been considered to be conspecific with ± 480 introduced from West Walsingham), West Hyporhamphus unifasciatus (Ranzani 1842) but is Walsingham Ponds (2,202 ± 178), Bartram’s Pond characterized by a more slender body, fewer pec- (1,793 ± 224, introduced from Lover’s Lake), and toral fin rays (usually 10), dorsal fin rays (usually Warwick Pond (436 ± 13). No estimates could be 14 or 15), and second arch gill rakers (usually 21- made for East Walsingham or Evan’s Pond. In addi- 23). Sequence data for 800 bp of mtDNA Cyt b tion, the annual reproductive cycle was described separates the Bermudan species from other species for the population inhabiting Mangrove Lake indi- of the genus by a minimum genetic distance of cating that males and females actively breed for 0.034 (Banford 2010). more than half the year, beginning in February and Fundulidae ending in September (Outerbridge et al. 2007b). Sequence divergence in the mitochondrial Cyt b Syngnathidae gene was assayed in four of the six extant Bermuda Cosmocampus albirostris (Kaup 1856), Whitenose killifish populations (Grady et al. 2001). Two Pipefish. Since the only two records of Cosmocam- divergent (4.6%) haplotypes were detected; one is pus albirostris supposedly from Bermuda were identical to the Georgia 2c haplotype of Fundulus without locality data, we (p. 189) decided that heteroclitus and is fixed in three eastern popula- additional voucher material was needed to confi- tions: Lover’s Lake (F. relictus), Mangrove Lake and dently include this pipefish in the known Bermuda Walsingham Pond (F. bermudae). The second is ichthyofauna. Russell Whayman found a specimen fixed and restricted to the westernmost population, (Fig. 22) near Northeast Breakers in August 2008 Evan’s Pond. Phylogenies and haplotype divergence and on a subsequent dive at the same spot found indicate at least two Bermuda colonizations, the two other individuals, confirming that this pipefish more recent involving transfer of the Georgia 2c does occur at least occasionally in Bermuda. haplotype. Discovery of killifish bones in peat Scorpaenidae deposits cored from Well Bay on Cooper’s Island in Pterois volitans (Linnaeus 1758), Red Lionfish. A St. David’s in a layer of organic sediments formed species introduced from the Indo-West Pacific and over 1,500 years ago (Outerbridge et al. 2006) pro- now widespread along the Atlantic coast of North vides physical evidence that at least one species of America (Whitfield et al. 2002; Meister et al. killifish was living in Bermuda 1,000 years before 2005; Morris et al. 2009), recently reported from humans arrived. Cytb sequences are uninformative several localities in the Bahamas (Snyder & Burgess as to the taxonomic status of the Bermuda 2007; Morris & Akins 2009), and other localities endemics F. bermudae and F. relictus, but support bordering on the Caribbean Sea (Schofield 2009, recognition of the Evan’s Pond population as an 2010). The first Bermuda record was based on a evolutionarily significant unit within the F. hetero- juvenile collected by Horace Landy in a Devon- clitus group. Genetic research on Bermuda’s killi- shire Bay tidepool, which was raised in his aquar- fish indicates that each pond has a genetically ium until it reached four inches long. This fish was unique population and that more endemic species entered in the 2001 Agricultural Exhibition and may be recognized in the future. Thus, it is imper- seized under suspicion of having been illegally

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imported into Bermuda (Sterrer 2002b; Whitfield of the opaque zones (interpreted as annuli) on et al. 2002). Spawning along the coast of south- transverse sections of the sagittal otolith gave an eastern United States probably supplied juvenile age estimate of 60 years. lionfish to Bermuda (Whitfield et al. 2002). Lion- Epinephelidae fish may cause deleterious changes in coral-reef Hyporthodus mystacinus (Poey 1852), Misty ecosystems by predation on native fishes and inver- Grouper. We follow Craig & Hastings (2007) and tebrates (Albins & Hixon 2008; Morris & Akins Craig et al. (2011) in placing this grouper in the 2009) so the fact that they are now abundant in genus Hyporthodus Gill rather than its traditional Bermuda raises concerns about their potential placement in Epinephelus Bloch. Two large speci- threat there. Morris et al. (2009) state that lionfish mens (152 and 157 cm TL) were caught in 2000- are established along the southeast U.S coast, 2001 by commercial fishermen using vertical long- Bermuda, and the Bahamas but presented no evi- line gear set in 220-270 m on the edge of the dence that lionfish are reproducing in Bermuda. Bermuda platform (Luckhurst & Dean 2009). The Bermuda Ministry of the Environment offers Counts of the opaque zones (interpreted as annuli) education and training seminars on lionfish so that on transverse sections of the sagittal otolith gave divers can apply for a license to net and spear lion- age estimates of 135 years for the 152-cm speci- fish (Morris et al. 2009; Dale 2009). mens, 150 years for the 157-cm specimen. These Brian Luckhurst had a very interesting session appear to be the oldest groupers reported to date with Chris Flook at the Bermuda Aquarium in the literature. Capture of these two groupers Museum and Zoo regarding lionfish. 1) All increased the documented maximum weight Bermuda specimens examined so far are identified (74.5 and 75.5 kg, respectively) for the species by as Pterois volitans based on fin-ray counts. 2) 50% over 20 kg. of females examined (N = 50) had ovaries with Hyporthodus niveatus (Valenciennes 1828), Snowy eggs, some apparently hydrated indicating immi- Grouper. We also follow Craig & Hastings (2007) nent spawning. Several very small juveniles (6-7 and Craig et al. (2011) in placing this grouper in cm) were collected suggesting a Bermuda origin. 3) the genus Hyporthodus Gill rather than its tradi- Eggs in Chris’ BAMZ tanks hatched in 4 days. He tional placement in Epinephelus Bloch. said that they grow very quickly and he believes Mycteroperca bonaci (Poey 1860), Black Grouper. A males mature in 6-7 months. 4) Chris has sampled spawning aggregation was located on the northeast stomach contents and has good evidence that lion- reef platform at a depth of about 30 m (Luckhurst fish are voracious predators in keeping with similar 2010). Diving observations revealed many similari- findings elsewhere. 5) In an effort to control pop- ties to observations of this species described at multi- ulation growth, Bermuda has licensed divers to species spawning aggregation sites in Belize. The spear lionfish and bring the specimens in to the spawning aggregations were observed at the warmest Aquarium. Currently about 130 divers are time of the year in Bermuda, June-August. licensed. Eradication is probably not possible but Paranthias furcifer (Valenciennes 1828), Creole-fish this policy may help control population growth. 6) or Barber. Hybridization of Paranthias furcifer and In October 2008, a lobster trap set on the edge (in Cephalopholis fulva was noted by us and treated in 30 fathoms) reportedly had 27 lionfish in one haul! detail by Bostrom et al. (2002). A recent molecular They may have entered the trap to eat other fishes phylogeny of groupers (Craig & Hastings 2007) caught in the trap. A fisherman reported another revealed that both species of Paranthias are nested anecdote to Chris: a vertical longline set on Chal- within Cephalopholis, implying that this species lenger Bank in 70 fathoms came up with 8 out of should be known as Cephalopholis furcifer. This new 10 hooks with a lionfish. The next day, same place, classification has been adopted by Page et al. (2013). 10 hooks, 10 lionfish! The fisherman had to move However, a major publication on groupers of the to another area because he could not catch any- world (Craig et al. 2011), as well as the Catalog of thing else there. Fishes (Eschmeyer 2013), reverted to the traditional Polyprionidae classification maintaining Paranthias as valid, “pend- Polyprion americanus (Bloch & Schneider 1801), ing additional study.” Thus, the generic placement Wreckfish. A 134-cm TL individual weighing 45.5 of this grouper is not fully resolved. kg was captured by commercial fishermen in Oct. Serranidae 1995 at 650 m (Luckhurst & Dean 2009). Counts Choranthias tenuis (Nichols 1920), Threadnose

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Bass. This species was moved from the genus Although we have been unable to examine his Anthias to their new genus Choranthias by Ander- material (these specimens cannot be located in the son & Heemstra (2012). Their account includes a institute’s collection, D. R. Robertson, pers. color description of a Bermuda specimen based on comm., Sept. 2012), Cervigón’s (1991) description a color transparency provided by the first author of and drawing (p. 403, Fig. 2) does not agree with this paper. the color photograph of Quattrini et al. (2004), Epinephelus guttatus (Linnaeus 1758), Red Hind. especially the less vertical orientation of the dark Luckhurst & Trott (2009) reported increases in size oblique bar in the spinous dorsal fin, in contrast to of Red Hind after protection by seasonally-closed that given in Bean’s (1912) original description spawning aggregation sites in Bermuda following “large black blotch on base [our italics] of spinous years of declining catches. dorsal extending upward a distance equal long Hypoplectrus puella (Cuvier 1828), Barred Hamlet. diameter of eye” and … “jet black spot behind base Only a single species of hamlet occurs in Bermuda. of pectoral fin.” Johnson & Smith-Vaniz (1987:50) Victor (2012) recently described a new species, noted that the pigment described by Bean was still Hypoplectrus floridae, from Florida that strongly evident on the holotype, including the black resembles H. puella, with which it had consistently “blotch in axil of pectoral fin.” However, the two been misidentified. The Florida species has a pair of most important discrepancies between Cervigón’s vertically aligned small dark spots at the base of the description (which agrees best with the holotype of caudal fin (absent in H. puella) and has a 3% mito- P. atrimanus) and P. incisus concern the shape of the chondrial DNA sequence-divergence from a large lacrimal (first infraorbital bone) and the relative homogenous clade of other Caribbean species of sizes of the two species. The lacrimal of the holo- Hypoplectrus including H. puella. Re-examination type is relatively deep with a broadly rounded pos- of specimens of the Bermuda hamlet reveals that terovental margin versus the slender and tapered they are the true H. puella and not H. floridae bone of P. incisus (see Johnson & Smith-Vaniz, which possibly could have been carried to the 1987, Figs 3c-d). Bean (1912) described the large island via the Gulf Stream. lacrimal of the type by stating “suborbital depth Parasphyraenops atrimanus Bean 1912, Bank Bass. 1/2 diameter of eye.” Cervigón’s drawing clearly This poorly known species, redescribed by Johnson shows a fish with a similarly broad lacrimal in con- & Smith-Vaniz (1987) based on the holotype and trast to photographs of P. incisus (Colin 1978; then only extant specimen, was questionably Quattrini et al. 2004) which show fish with narrow treated by us (p. 216) as a Bermuda endemic. How- suborbital depths. Cervigón’s three specimens were ever, Heemstra et al. (2003: 1327) gave the distrib- reported to be 64-72.5 mm SL, all exceeding the ution of P. atrimanus as Bermuda and Venezuela. A largest known individual of P. incisus (see above). second species of small planktivorus serranid, origi- Rypticus carpenteri Baldwin & Weigt 2012, Slope nally described as Serranus incisus by Colin (1978), Soapfish. This new species has been confused with was transferred to Parasphyraenops by Johnson & the Spotted Soapfish, R. subbifrenatus, with which Smith-Vaniz (1987). The reported Venezuelan dis- it shares a very similar pattern of dark spots. Both tribution for Parasphyraenops atrimanus was based species are broadly distributed throughout the on Heemstra’s conclusion (pers. comm., March, Caribbean and occur sympatrically at many locali- 1995) that Cervigón’s (1991:403) description and ties. We erroneously identified (p. 218) the only figure of Serranus incisus actually apply to P. atri- two collections of Bermuda soapfish that have con- manus, an opinion with which we agree. spicuous dark spots as R. subbifrenatus; however, Colin (1978) described Serranus incisus from re-examination of the two ANSP specimens by material collected from Jamaica and Puerto Rica Baldwin & Weigt (2012) has shown them to actu- and observed but did not collect specimens off ally be R. carpenteri. The two historical FMNH Curaçao that he believed to be this species. Subse- specimens are badly faded and, although probably quently, Quattrini et al. (2004) recorded it from R. carpenteri, cannot be positively identified. deep reefs off North Carolina, reported it from the Rypticus bistrispinus (Mitchill 1818), Freckled Bahamas and Turks and Caicos (R. G. Gilmore, Soapfish. In our synonymy of Rypticus saponaceus pers. comm.), and documented the life colors (p. 218) we listed Bean’s (1906:55) report of a sin- (their Plate 1b) of a 55 mm SL specimen, the gle specimen (FMNH 5295) of R. bistrispinus from largest yet recorded of this diminutive planktivore. Nonsuch Island, Bermuda as a misidentification.

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However, re-examination of the specimen by both Atlantic Bumper. Only three positively identified us and Guimarães (1999) has confirmed the accu- specimens of bumper from Bermuda existed when racy of the original identification. This in the only we wrote the book (p. 239) but recently there was confirmed record of the species from Bermuda. an influx of small bumper into Bermudian waters Malacanthidae where some were netted by Kevin Winter and Caulolatilus bermudensis Dooley 1981, Bermuda Brian Lines on April 30, 2006 while hauling bait at Tilefish. The life colors of this Bermuda endemic Foot of the Lane. Brian Luckhurst thinks these (Fig. 23) were unknown when the species was orig- may have arrived on a Gulf Stream eddy. Repre- inally described and this is the first photograph of sentative specimens have been preserved in the a fresh specimen. BAMZ collections as 2006-238-004. Caulolatilus dooleyi Berry 1978, Bahama Tilefish. Bramidae This tilefish was originally described from a single Pteraclis carolinus (Valenciennes in Cuvier & specimen from the Bahamas (Berry 1978). Valenciennes 1833), Atlantic Fanfish. Two speci- Recently we received a photograph (Fig. 24) of mens (Fig. 26) were found in the stomach of a Yel- what appears to be this species (Dooley, pers. com.) lowfin Tuna off the Banks in 2004 (communicated but because the specimen was not retained and no to us by Chris Flook, BAMZ). This record adds meristic or morphometric data were taken, the another species to the three bramids known from identification should be considered tentative until Bermuda. more specimens become available. Gerreidae Rachycentridae Gerres cinereus (Walbaum 1792), Yellowfin Rachycentron canadum (Linnaeus 1766), Cobia. A Mojarra. Because there was only one confirmed rare fish in Bermuda known from (p. 233) some record of this species from Bermuda (p. 260), we early Bermuda records: a museum specimen caught considered it to be a non-established waif species. in 1957, and a 124-cm TL specimen taken at Wat- However, REEF volunteers have now recorded the ford Bridge, Somerset in June 1985; thus, a recent species from Harrington Sound, Flatts Inlet, Wals- capture is of interest. A three-foot cobia was caught ingham Pond, Bailey’s Bay, Evans Bay, Shelly Bay, and photographed by Denis Kerr off the South Parsons Bay, Grotto Bay, Tom Woods Bay, Elys Shore in June 2010 (Arandjelovic 2010). Harbour mangroves, Penhurst Park, etc. (Judie Carangidae Clee, pers. com., 12 May 2013) indicating that the In the key to the jacks in our book (p. 235), there Yellowfin Mojarra (Fig. 27) is undoubtedly an is an error in couplet 13. It should read gill rakers established Bermuda species. 10-14 upper and 25-28 lower versus gill rakers 6- Haemulidae 9 upper and 16-21 lower instead of lower and Haemulon macrostomum Günther 1859, Spanish total. Grunt. Although there were historical reports of Caranx bartholomaei Cuvier 1833, Yellow Jack. In Spanish grunts frequently being brought into 1999 (p. 237), we believed that all the Bermuda Hamilton by fishermen, who caught them in deep specimens of this species were juveniles 62-82 mm water near the outer reefs, we incorrectly con- FL, suggesting a non-resident waif occurrence. cluded (p. 263) that these and other literature However, we overlooked a 350-mm FL specimen, records were based on misidentifications because ANSP 177887 taken off Kitchen buoy by P. C. Bermuda fisheries workers were unfamiliar with Heemstra and J. Burnett-Herkes in June 1973 so the species and we were unable to locate any cor- adults do at least occasionally reach Bermuda. rectly identified museum voucher specimens. Caranx hippos (Linnaeus 1766), Crevalle Jack. We However, Drummond-Hay’s unmistakable water- reidentified the only specimens identified as color (Fig. 10) documents at least the historical Caranx hippos as C. latus and concluded, as did presence of the species in Bermuda. Smith-Vaniz & Carpenter (2007), that the Crevalle Inermiidae Jack is a continental species that does not occur in Haemulon vittatum (Poey 1860), Boga. This fam- Bermuda. However, we have received an underwa- ily is now considered a synonym of Haemulidae ter photograph (Fig. 25) from Lisa Greene, taken and the species formerly known as Inermia vittata at Aristo Wreck in 2009, that proves that Crevalle is a member of the genus Haemulon, based on Jack do occur, at least occasionally, in Bermuda. mitochondrial and nuclear genetic data (Rocha et Chloroscombrus chrysurus (Linnaeus 1766), al. 2008; Sanciangco et al. 2011).

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Fig. 3. Holocentrus rufus, Longspine Squirrelfish, Drum- mond-Hay no. 41, "Squirrel fish," natural size, original drawing about 163 mm TL, Bermuda, 21 October 1847. Fig. 6. Lutjanus synagris, Lane Snapper, Drummond-Hay Courtesy of Smithsonian Institution, no. P12956. no. 12, “White water snapper,” 2/3 natural size, original drawing 183 mm TL, Bermuda, 12 March 1849. Courtesy of Smithsonian Institution, no. P11884.

Fig. 7. Eucinostomus gula, Silver Jenny, Drummond-Hay no. 18, “Shad,” natural size, 72 mm TL, Bermuda, Sep- tember 1850. Courtesy of Smithsonian Institution, no. Fig. 4. Hippocampus erectus, Lined Seahorse, Drummond- P10700. Hay no. 89, “Sea horse,” natural size, original of the larger drawing ca. 100 mm TL, Bermuda, October 1847. Cour- tesy of Smithsonian Institution, no. P15911.

Fig. 5. Mycteroperca tigris, Tiger Grouper, Drummond- Fig. 8. Haemulon album, Margate, Drummond-Hay no. Hay no. 4, “Gag,” 1/2 natural size, original drawing 270 24, “Margate fish,” 1/2 natural size, original drawing 178 mm TL, Bermuda, November 1849. Courtesy of Smith- mm TL, Bermuda, 19 June 1847. Courtesy of Smithson- sonian Institution, no. P9679. ian Institution, no. P12039.

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Fig. 9. Haemulon flavolineatum, French Grunt, Drum- Fig. 12. Kyphosus vaigiensis, Yellow Chub, Drummond- mond-Hay no. 20, “Large scale yellow grunt,” natural size, Hay no. 28, “Chub,” natural size, original drawing 201 original drawing 176 mm TL, Bermuda, October 1850. mm TL, Bermuda, 14 September 1850. Courtesy of Courtesy of Smithsonian Institution, no. P12038. Smithsonian Institution, no. P14490.

Fig. 10. Haemulon macrostomum, Spanish Grunt, Drum- Fig. 13. Bathygobius lacertus, Spotted Frillfin, Drummond- mond-Hay no. 21, “Black grunt,” 2/3 natural size, original Hay no. 60, “Goby”, natural size, original drawing 102 drawing 186 mm TL, Bermuda, 20 May 1851. Courtesy mm TL, Bermuda, 15 March 1851. Courtesy of Smith- of Smithsonian Institution, no. P12036. sonian Institution, no. P10723.

Fig. 11. Haemulon sciurus, Bluestripe Grunt, Drummond- Fig. 14. Cantherines macrocerus, Whitespotted Filefish, Hay no. 19, “Small scaled yellow grunt,” common size, Drummond-Hay no. 96, “Ocean turbot, 1/2 natural size, original drawing 188 mm TL, Bermuda, July 1849. Cour- original drawing 224 mm TL, Bermuda, February 1851. tesy of Smithsonian Institution, no. P12037. Courtesy of Smithsonian Institution, no. P2285.

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Fig. 15. Canthigaster rostrata, Sharpnose Puffer, Drum- mond-Hay no. 93, “Puff fish,” natural size, original draw- ing 57 mm TL, Bermuda, no date. Courtesy of Smithson- Fig. 17. Dasyatis centroura, Roughtail Stingray, Bermuda. ian Institution, no. P2285. Photo by G. Mayer.

Fig. 18. Ophichthus ophis, Spotted Snake Eel, Clearwater Beach, Bermuda. Photo by S. R. Smith.

Fig.16. Masturus lanceolatus, Sharptail Mola, Drummond- Hay unnumbered, “Short sun-fish,” 4 ft. 3 inches long, Bermuda, 31 May 1851. Courtesy of Smithsonian Institu- tion, no. P11073.

Fig. 20. Carapus bermudensis, Pearlfish, Bermuda. Photo Fig. 19. Lampris guttatus, Opah, Bermuda. Photo from D. by W. Sterrer. Soares.

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Fig. 25. Caranx hippos, Crevalle Jack, Astro Reef, Bermuda. Photo courtesy Lisa Greene, Bermuda Natural History Museum. Fig. 21. Antennarius scaber, Splitfin Frogfish, Bermuda. Photo by K. Holzer.

Fig. 22. Cosmocampus albirostris, Whitenose Pipefish, Fig. 26. Pteraclis carolinus, Atlantic Fanfish, from the stom- Bermuda. Photo by R. Lucas. ach of a Yellowfin Tuna off the Banks, Bermuda in 2004. Photo by S. Cabral, Jr.

Fig. 23. Caulolatilus bermudensis, Bermuda Tilefish, Bermuda. Photo courtesy of Bermuda Department of Environmental Conservation. Fig. 27. Gerres cinereus, Yellowfish Mojarra, Bermuda. Photo by L. Franks.

Fig. 24. Caulolatilus dooleyi, Bahama Tilefish, Bermuda. Photo courtesy of Bermuda Department of Environmental Fig. 28. Cynoscion regalis, Weakfish, Bermuda. Photo by Conservation. T. Adderly.

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Fig. 33. Halichoeres garnoti, Yellowhead Wrasse, with typi- cal “decoratus” colour pattern (see text discussion). Photo by R. Lucas.

Fig. 29. Pomacanthus paru, French Anglefish. Bermuda. Photo by J. Burville.

Fig. 34. Halichoeres poeyi, Blackear Wrasse, Bermuda. Photo by L. Franks.

Fig. 30. Chromis bermudae, Yellowfin Chromis, Bermuda. Photo by A. Marquart.

Fig. 31. Chromis flavicauda, Cobalt Chromis, Brazil. Photo by A. de Luca, Jr. Fig. 35. Masturus lanceolatus, Sharptail Mola, Horseshoe Beach, Bermuda. Photo by A. Davis.

Fig. 32. Clepticus sp., Bermuda Creole Wrasse, Bermuda. Fig. 36. Ranzania laevis, Slender Mola. Photo by D. Skin- Photo by R. Lucas. ner.

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Sciaenidae L. Gasparini and others, based on a large series of Cynoscion regalis (Bloch & Schneider 1801), fresh C. flavicauda obtained during recent surveys Weakfish. A Weakfish (Fig. 28) was caught on Oct. of deep reefs off northeastern Brazil, reveal that 16, 2001 by Thomas Adderly on the North Shore they consistently differ from the Bermuda Chromis inside the shipping channel off of Bailey’s Bay in in having the caudal peduncle and spinous dorsal 40-60 feet of water. This is a new record for almost entirely blue versus a broad diagonal band Bermuda. The specimen has been catalogued as of yellow on the caudal peduncle extending BAMZ 2001-197-024. anterodorsally to the rear of the dorsal fin and at Pomacanthidae least the distal third of the spinous dorsal fin bright Pomacanthus paru (Bloch 1787), French yellow. Bermuda fish also have a more extensive Angelfish. We reported this angelfish as historically area of yellow in the anal fin. introduced to Bermuda with the most recent Considering the color pattern differences between recorded sighting in October 1968. However, Rus- the Bermuda and Brazilian Chromis (Figs. 30-31), sell Whayman sent us photographs of a pair of their widely disjunct distributions, and recent pub- adults he observed in Bermuda on August 17 lications indicating that the level of endemism 2006. They have been periodically seen by several of the Brazilian reef fishes is much greater than divers off East End and one of the pair was pho- previously thought (Menezes et al. 2003; Moura tographed (Fig. 29) by John Burville. Whether the & Sazima 2003; Rocha 2003), we now recognize species is reproducing in Bermuda is uncertain. Chromis bermudae as a valid Bermudian endemic. Kyphosidae Labridae Species of Kyphosus are difficult to identify and a Clepticus sp., Bermuda Creole Wrasse. We recent family revision by Knudsen & Clements recently became aware of two color photographs, (2013) has revealed that in addition to the Bermuda one of a school and the other of a single individual Chub, Kyphosus sectatrix, two other chubs occur in (the latter here reproduced as Fig. 32), of wrasses Bermuda, both with unfamiliar names. from Bermuda (Lucas 2012) identified as Clepticus Kyphosus vaigiensis (Quoy & Gaimard 1825), Yel- parrae. Terminal phase adults are distinctive in hav- low Chub. We recorded this chub (p. 278) as K. ing very elongate outer caudal-fin rays, mostly incisor (Cuvier 1831), a species then considered to solid blue body and fins and a bright yellow snout. be restricted to the Atlantic Ocean. Knudsen & We predict that subsequent research will show Clements (2013) have determined that it actually these fish to be another Bermuda endemic. All occurs in both the Atlantic and Indo-Pacific records of Clepticus parrae from Bermuda appar- regions and was first described by Quoy & ently are based on misidentifications of this unde- Gaimard (1825) from Indonesia. scribed Clepticus. Heiser et al. (2000) provide the Kyphosus bigibbus Lacepède 1801, Brown Chub. most recent review of Clepticus species. Previously confused with K. sectatrix, Knudsen & Halichoeres garnoti (Valenciennes 1839), Yellow- Clements (2013) recorded this species from both head Wrasse, Fig. 33. We noted (p. 293) that ter- sides of the Atlantic and the Indo-Pacific region, and minal males of H. garnoti from Bermuda (our listed the following three specimens from Bermuda: colour Pl. 12, Figs 78-79) “typically differ from ZMB 7866 (100 mm SL), ZMB 7997 (186 mm non-Bermuda individuals in having a dorsum that SL) and ZMUC journ. 256 (150 mm SL). is bright orange to vermilion instead of being yel- Pomacentridae lowish or greenish posterior to a diagonal cross- Chromis bermudae Nichols 1920, Bermuda band at the origin of the first segmented dorsal-fin Chromis. We treated (p. 284) this colorful dam- ray,” and that should subsequent studies reveal selfish as a junior synonym of C. flavicauda (Gün- additional differences, the name H. decoratus ther 1880), described from northeastern Brazil, (Bean) is available for the Bermuda form. following Smith-Vaniz & Emery (1980) who had In a study of mitochondrial DNA variation in only the badly faded Brazilian holotype for com- western Atlantic Halichoeres, Rocha (2004) deter- parison. Although recognizing these two nominal mined that H. garnoti had no significant popula- species as conspecific, Moura (1995) noted that tion structure across its entire range from Bermuda Brazilian specimens have less yellow coloration on to Venezuela; the Bermuda population shares 12 of the caudal peduncle, dorsal and anal fins. Observa- its 20 haplotypes with fish at one or more tions and photographs provided by L. A. Rocha, J. Caribbean locations, and haplotypes restricted to

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Bermuda were nested within the Caribbean lin- The species that we recorded (p. 312) as Eleotris eage. Rocha reported that Halichoeres garnoti does pisonis is actually Eleotris perniger (Cope 1871). not occur in the northern Gulf of Mexico Pezold & Cage (2002) have determined that E. (although there are records from deep reefs in the pisonis is a continental South American species northwest Gulf of Mexico) or in any coastal loca- while the insular Caribbean basin species is E. tion north of Florida and presumably is a strictly perniger. tropical species. During the last glacial maximum Gobiidae (25,000-15,000 yr B.P.) temperatures at Bermuda In our book we reported only two Bathygobius were much lower than present (Sachs & Lehman species from Bermuda. A recent review of western 1999), thus it is probable that the Bermuda popu- Atlantic Bathygobius by Tornabene et al. (2010) lation is relatively young and although the color requires two additions to the species composition differences likely have a genetic basis, they are not of the genus in Bermuda. The color pattern con- detectable in neutral genes, such as cytochrome b, sisting of a double row of diagonal, dark blotches because of insufficient time for lineage sorting. on the flanks clearly identifies Drummond-Hay’s Halichoeres poeyi (Steindachner 1867), Blackear “Goby” watercolor (Fig. 13) as that of Bathygobius Wrasse. The first record of this wrasse from lacertus (Poey 1860). Apparently, at least four Bermuda was based on a solitary, initial phase, species of Bathygobius occur in Bermuda (L. Torn- individual observed at Baileys Bay on 25 Septem- abene and C. Baldwin, pers. com.): B. soporator, B. ber 2002. While volunteering with a biodiversity curacao, B. lacertus, and B. antilliensis Tornabene, team conducting seagrass transects, Judie Clee Baldwin, & Pezold 2010. Tornabene et al. (2010) (pers. com.) sighted it amongst a school of Slippery re-identified the holotype of Bathygobius soporator Dicks, Halichoeres bivittatus (Bloch) and kept an sextaneus Ginsburg 1947 as probably a specimen of eye on it for an extended period of time. She was B. lacertus and reported that additional museum very familiar with the Blackear Wrasse having seen specimens (without catalog numbers) of this them many times in the Caribbean and was species were examined from Bermuda (see also absolutely positive of the identification, based on Tornabene et al. 2010, Fig. 13). The specimens its very pea-green colour and distinctive ear mark. (ANSP, UMMZ) that we previously identified as Lydia Franks observed and photographed (Fig. 34) B. curacao are apparently correctly identified. a second initial phase individual between Flatts and However, material that we identified as B. soporator Gibbet Island on September 4, 2012. Because contains all four species. One lot (USNM 178014) these are the only records of this wrasse in collected by William Beebe in 1929 contains a few Bermuda, we consider the two observations of sin- B. soporator and multiple specimens of B. curacao gle individuals to be examples of non-established (L. Tornabene, pers. com.). ANSP 32677 is re- waif occurrences. identified as B. lacertus and ANSP 14287 as prob- Three species of parrotfishes were observed in a ably B. antilliensis (C. Baldwin, pers. com.). multispecies spawning aggregation at the same UMMZ material originally identified as B. sopora- time and location just outside the breaking reef tor was reidentified (C. Baldwin, pers. com.) as line along the southwest coast of Bermuda about either B. antilliensis (UMMZ 172393) or B. cura- 500 m from the shoreline in June and July 2003 cao (UMMZ 176373, 1764562). (Luckhurst 2011). The species were the Redfin Parrotfish Sparisoma rubripinne (46 spawning Key to Bermuda Bathygobius events observed), Queen Parrotfish Scarus vetula (based on Tornabene et al. 2010) (11), and Stoplight Parrotfish, Sparisoma viride (3). 1a. First dorsal fin with a broad vertical or slightly Uranoscopidae diagonal dark bar...... B. soporator Xenocephalus egregius (Jordan & Thompson 1b. First dorsal fin with a longitudinal pattern of 1905), Freckled Stargazer. We follow Springer & pigment, either a single broad stripe or 1-4 nar- Bauchot (1994) and Eschmeyer (2013) in assign- row stripes ...... 2 ing this stargazer to the genus Xenocephalus Kaup 2a. Ventral portion of trunk with two rows of 6-7 1858 instead of its traditional classification in dark blotches, the blotches in the two rows off- Gnathagnus Gill 1861, a junior synonym. set and reminiscent of the dark squares on a Eleotridae checker-board; lateral scale rows 38-42 ...... Eleotris perniger (Cope 1871), Spinycheek Sleeper...... B. lacertus

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2b. Ventral portion of trunk variously pigmented, terns, but there are no confirmed Bermuda collec- with or without blotches, but never with two tions of the latter species. Presence of a third offset rows of blotches; lateral scale rows 31-42 species, C. bol, is based mostly on genetic charac- ...... 3 ters making identification of it difficult. We pre- 3a. Ventral portion of trunk with two rows of sent the following revised key to Bermuda markings, the uppermost with 7-8 dark Coryphopterus. blotches beginning beneath the pectoral fin and terminating just before basicaudal mark- Key to Bermuda Coryphopterus (modified, in ing; lower row with 3 (rarely 4) spots and ter- part, from Baldwin et al. 2009) minating anterior to anal fin origin; lateral 1a. Pelvic fins broadly united ...... 2 scale rows 38-42; pectoral fin rays 18-21 ...... 1b. Pelvic fins completely separate or united ...... B. antilliensis basally by narrow membrane ...... 4 3b. Trunk typically with no diagnostic pigment 2a. Body usually pale, pigment primarily compris- pattern (sometimes 6-7 diffuse dark blotches of ing three rows of markings on side of body; varying size along body); lateral scale rows 31- lower row comprising small, mostly vertically 36; pectoral fin rays 15-17...... B. curacao elongate markings, some of which may be cres- cent shaped or some part of an X-shape but Coryphopterus tortugae (Jordan 1904), Sand Goby. rarely well defined X’s; if X-shaped markings Although Garzón & Acero (1990) resurrected this present, their height is considerably shorter species from synonymy with C. glaucofraenum Gill than eye diameter; pigment marking above 1863 based on color pattern (basicaudal marking a opercle usually a triangle, and basicaudal pig- bar instead of two colon-like spots) and relative ment usually a central bar...... C. tortugae body depth, we (p. 315) felt that the color pattern 2b. Body heavily pigmented or pale but without difference was not sufficient to recognize two dis- vertically elongate or crescent-shaped markings tinct species in Bermuda. However, Victor (2008) in ventral row of pigment on side of body; and Baldwin et al. (2009) used morphological and height of X-shaped markings, if present, three- molecular characters to successfully differentiate quarters of or equal to diameter of eye; pig- the species. Re-examination of most of our mater- ment marking above opercle triangular, ial showed both species to be common but usually rounded, or with two well-defined peaks; basi- not occurring together. Of 41 stations shown on caudal pigment comprising two separate spots, the map in Fig. 128, only five stations (81-4, 81-5, two spots connected by a line of pigment and 81-6, 81-10 and 81-32) contained specimens of resembling a dumbbell, a central bar, or a C- both species. Half of the other stations (18) con- shaped...... 3 tained only C. glaucofraenum, and half (18) con- 3a. Pigment on pectoral-fin base variable but tained only C. tortugae. One specimen from Three always with dark spot or rectangular-shaped Hills Shoals that was originally identified as C. blotch ventrally (may be associated with bright glaucofraenum has been reidentified by B. Victor, yellow pigment in life); one or two additional based on genetic evidence, as C. bol Victor 2008 bars, blotches, or concentrations of pigment (A. Bentley, pers. comm.). However, Baldwin et al. sometimes present dorsally; three rows of dark (2009: 129) consider C. bol to be a synonym of C. markings on side of body, some in lower row venezuelae Cervigón 1966. Additional research is large, X-shaped markings in heavily pigmented needed to resolve this issue. specimens, small, circular blotches in paler We do not list the stations where each of these specimens; pigment markings above the oper- two species of Coryphopterus were collected sepa- cle triangular or round ...... C. venezuelae rately but for those who want that information it 3b. Pectoral-fin base rarely with prominent dark can be obtained by accessing the ANSP ichthyol- marking ventrally, although melanophores may ogy collection database at http://data.acnatsci.org/ form one to three light to moderate concentra- biodiversity_databases/fish.php. Coryphopterus tions on base; body with three rows of dark glau cofraenum and C. venezuelae Cervigón 1966 markings, most of those in the lower row large, (originally described as a subspecies of C. glau- distinctive X-shaped markings; pigment mark- cofraenum) co-occur throughout much of their ing above opercle usually with two well-defined respective ranges and have very similar color pat- peaks ...... C. glaucofraenum

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4a. Anus situated at or near center of black ring; in 1980 that we recorded in our book and Andrew anterior interorbital region of head with one Card recently caught another one off Argus Bank median pore ...... C. personatus on 1 Feb. 2012, which weighted 920 lbs even with 4b. Anus not situated at center of black area but part of its caudal peduncle eaten by sharks nearer posterior margin; anterior interorbital (Bernews 2012). region of head with two pores, each nearer Istiophoridae orbital rim than fellow pore ...... C. hyalinus Collette et al. (2006) revised the family and rec- ommended several changes in nomenclature of Gnatholepis cauerensis (Bleeker 1853), Goldspot billfishes affecting one species reported from Goby. We recorded (p. 316) this common and Bermuda. The White Marlin Tetrapturus albidus widely distributed Caribbean goby as Gnatholepis (p. 336), along with the Indo-Pacific Striped Mar- thompsoni Jordan 1904 because, as the only lin, have been moved to the genus Kajikia Hirasaka Atlantic representative of the genus Gnatholepis, and Nakamura 1947 so its name is now Kajikia application of the scientific name had not been albida (Poey 1860). questioned. In a study of mitochondrial DNA vari- Makaira nigricans Lacepède 1802, Blue Marlin. ation in populations of Gnatholepis, Rocha et al. While fishing offshore in July 2006, Ian Card was (2005) presented data indicating that a single speared in the chest and knocked overboard by a species had colonized the Atlantic from the Indian blue marlin estimated to be about 363 kg and 4.3 Ocean during the late Pleistocene. Subsequently, m long. The marlin had just been hooked when it Randall & Greenfield (2007) redescribed Gobius suddenly leaped out of the water and impaled Mr. cauerensis Bleeker, 1853, discussed why the name Card just below his collar bone. The bill just used by Rocha et al. (op. cit.) was incorrect and missed one of his main arteries so the wound easily concluded that Gnatholepis cauerensis (Bleeker) is a could have been fatal. senior synonym of G. thompsoni. Based on gonad histology, 70% of 11 female Blue In their recent revision of the genus Gnatholepis, Marlin caught in July in Bermuda waters were Larson & Buckle (2012) treat Gnatholepis thomp- actively spawning or in spawning condition (Luck- soni and G. cauerensis as allopatric sister-species. hurst et al. 2006). These observations confirm that However, they stated that fish from South Africa, active spawning of Blue Marlin occurs in Bermuda which they identify as G. cauerensis, “look almost waters in July extending the known spawning identical in colour pattern to G. thompsoni” as do range of the species in the western North Atlantic. certain others from Indonesia and Malaysia, and Also, a juvenile (256 mm lower jaw fork length), fish from Atol das Rocas, Brazil “resemble G. estimated to be 42 days old, was dipnetted on cauerensis in colouring,” as do ... photographs of Challenger Bank on Sept. 2, 1994. fish from San Salvador which “show similar- Molidae coloured fish.” We acknowledge the thoroughness We recorded (p. 358) two species of Molidae from of their revision, but note that all the meristic and Bermuda, Mola mola and Ranzania laevis. However, other morphological characters purported to sepa- the first report (Goode 1877) of Mola mola (as M. rate these two “species” have broadly over-lapping rotunda) from Bermuda was actually based on values. We therefore disagree with their choice of misidentification of a third species, the Sharptail nomenclature, which seems to have been strongly Mola, Masturus lanceolatus (Liénard 1840). This biased by geography. became obvious when a recently discovered Drum- Acanthuridae mond-Hay sketch (Fig. 16) included collection data Based on color pattern and genetic differences, identical to that reported by Goode. This circum- Bernal & Rocha (2011) showed that the Bermuda global species differs from other molas in having a and northwestern Atlantic population of the Ocean protruding pseudocaudal fin (clavicle) and a hori- Surgeon that we referred to (p. 321) as Acanthurus zontal mouth. Andrew Davis found a second speci- bahianus Castelnau 1855 is actually a distinct men of the Sharptail Mola floating, alive, in shallow species, Acanthurus tractus Poey 1860. The South water at Horseshoe Beach, Southampton on May Atlantic species retains the name A. bahianus. 27, 2001. The 407-mm SL specimen (Fig. 35) is Scombridae catalogued as BAMZ 2001-194-001. Thunnus thynnus (Linnaeus 1758), Atlantic Ranzania laevis (Pennant 1776), Slender Mola. Bluefin Tuna. Alan Card caught the Bluefin Tuna The only previous Bermuda specimen of this

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molid was one obtained from the stomach of a may have colonized Bermuda during a previous Wahoo caught on Argus Bank in 1995. Recently interglacial warming period 120,000-130,000 two additional specimens of this rarely seen species years ago. (Fig. 36) were discovered on the East End: Judie Grady et al. (2001) also found molecular evidence Clee found one on the South Shore of Coopers of multiple colonization events in killifishes of the Island in St. David’s and Chris Smith found genus Fundulus. However, we reported (p. 179) another one the same day (April 7, 2002) at Clear- that age estimates based on radiometric dating of water Beach. The two specimens have been cata- peat deposits for Mangrove Lake and other isolated logued as BAMZ 2002-203-001 (445 mm TL) and Bermuda inland marshes, range from 4-8+ thou- BAMZ 2002-203-002 (430 mm TL). sand years. Prior to the rapid rise in sea levels Endemism and Phylogeography: The description beginning about 7,000 years BP Fundulus species of the Bermuda population of Hyporhamphus as a would have been unable to surmount the steep new species, recognition of Chromis bermudae flanks of the Bermuda platform. (Pomacentridae) as a valid species, and removal of Parasphyraenops atrimanus (Serranidae) from the list ACKNOWLEDGEMENTS of Bermuda endemics, changes the total number of Many colleagues have called our attention to addi- Bermuda endemic fishes to seven species (eight if tional records, references, additional Drummond- the Bermuda Clepticus proves to be another Hay illustrations, name changes, errors in our 1999 endemic, as we believe), excluding from considera- book, or helped with identifications. We especially tion several land-locked species of Fundulus. It is thank Brian Luckhurst, formerly of Bermuda Fish- difficult to determine if some species (including eries and co-author of our original book, and the several discussed above) are established in Bermuda staff and associates of the Bermuda Aquarium, Nat- but with very low population numbers or they rep- ural History Museum, and Zoo who have continued resent rare waif occurrences. Regardless of how to provide us with useful information about these species are categorized or the definition used Bermuda fishes as did Carole Baldwin, Judie Clee, to determine which families (or species) should be Daniel Cohen, Chris Flook, Lisa Greene, Karsten treated as “shorefishes,” the percentage of nearshore Hartel, Luiz Rocha, John McCosker, John Bermuda endemic fishes is less than 3 percent. McEachran, Susan Mochel, Mark Outerbridge, Lisa Sequence analysis of the mitochondrial control Palmer, Ross Robertson, Struan Smith, Wolfgang region and the nuclear ITS1 region was performed Sterrer, Luke Tornabene and no doubt others as well. to investigate some questions about Bermudian For making important color photographs available phylogeography (Johnson 2003). Seven species of for our use we also thank Thomas Adderly, John common shorefishes were analyzed, eight individu- Burville, Andrew Davis, Armando de Luca, Jr., Lydia als from Bermuda and eight others from the south- Franks, Lisa Greene, Kimberly Holzer, Raphael eastern United States. Haplotype diversities of the Macieira, Alan Marquart, Gunar Mayer, Ron Lucas, control region and gene diversities of ITS1 in Jo Pitt, David Soares, David Skinner, Struan R. Bermudian populations were relatively high, while Smith and Wolfgang Sterrer. This is contribution # mean nucleotide sequence diversities of the control 212, Bermuda Biodiversity Project (BBP), Bermuda region and ITS1 were relatively low. This is indica- Aquarium, Natural History Museum and Zoo, tive of rapid population growth following a period Department of Conservation Services. of low effective population size or a founder event. Four species, Haemulon flavolineatum, Holocentrus REFERENCES adscensionis, Lagodon rhomboides, and Lutjanus ALBINS, M. A. & HIXON, M. A. 2008. Invasive Indo- griseus, have mitochondrial control region and Pacific lionfish Pterois volitans reduce recruitment of ITS1 divergence values corresponding to mean Atlantic coral-reef fishes. Marine Ecology Progress Series divergence times of 2,000 to 21,500 years which 367: 233-238. suggests colonization following the last glacial ANDERSON, W. D., JR. & HEEMSTRA, P. C. 2012. Review of period 18,000 years ago. Three species, Haemulon Atlantic and eastern Pacific anthiine fishes (Teleostei: Per- ciformes: Serranidae), with descriptions of two new gen- aurolineatum, H. sciurus, and Holocentrus rufus era. Transactions of the American Philosophical Society 102 have mitochondrial control region and ITS1 diver- (2): 1-173. gence values corresponding to mean divergence ARANDJELOVIC, N. 2010. Rare cobia fish caught off the times of 38,000 to 214,500 years. These species South Shore. The Royal Gazette, Tuesday, June 29, 2010.

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ARNOLD, R. J. & PIETSCH, T. W. 2012. Evolutionary history COLIN, P. L. 1978. Serranus incisus, new species from the of frogfishes (Teleostei: Lophiiformes: Antennariidae): a Caribbean Sea (Pisces: Serranidae). Proceedings of the Bio- molecular approach. Molecular Phylogenetics and Evolution logical Society of Washington 19 (1): 191-196. 62: 117-129. [Available online 2 Oct. 2011; publisher CRAIG, M. T. & HASTINGS, P. A. 2007. A molecular phy- listed hard copy as 2012]. logeny of the groupers of the subfamily Epinephelinae BALDWIN, C. C. & WEIGT, L. A. 2012. A new species of (Serranidae) with a revised classification of the Epineph- soapfish (Teleostei: Serranidae: Rypticus), with redescrip- elini. Ichthyological Research 54 (1): 1-17. tion of R. subbifrenatus and comments on the use of DNA CRAIG, M. T., SADOVY DE MITCHESON, Y. J. & HEEMSTRA, barcoding in systematic studies. Copeia 2012 (1): 23-36. P. C., eds. (2011). Groupers of the world. A field and mar- BALDWIN, C. C., WEIGT, L. A., SMITH, D. G. & MOUNTS, ket guide. NISC (Pty) Ltd. Grahamstown, South Africa, J. H. 2009. Reconciling genetic lineages with species in 356 pp. + A: 1-47. western Atlantic Coryphopterus (Teleostei: Gobiidae). In: DALE, A. 2009. It’s open season on the lionfish. The Royal Proceedings of the Smithsonian Marine Science Symposium, Gazette, Feb. 19, 2009. www.royalgazette.com. ed. M. A. Lang, I. G. Macintyre, and K. Rützler. Smith- ESCHMEYER, W. N. (ed.) 2013. Catalog of Fishes. Electronic sonian Contributions in Marine Science 38: 111-149. version http://www.calacademy.org/ research/ichthyol- BANFORD, H. M. 2010. Hyporhamphus collettei, a new ogy/catalog [accessed 30 April 2013]. species of inshore halfbeak (Hemiramphidae) endemic to FRABLE, B. J., BALDWIN, C. C., LUTHER, B. M. & WEIGT, Bermuda, with comments on the biogeography of the L. A. 2013. A new species of western Atlantic lizardfish Hyporhamphus unifasciatus species group. 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Vol. 2: Bony fishes part 1 (Acipenseridae to tion. Fundación Científica Los Roques, Caracas, Grammatidae). FAO Species Identification Guide for Fishery Venezuela, vol. 1, 425 pp. Purposes and American Society of Ichthyologists and Herpetol- COHEN, D. M., HENSLEY, D. A. & KIMMEL, J. J. III. 1991. ogists Special Publication 5: 1071-1085. The deepsea ophidiid fish genus Lamprogrammus, a senior HARRISON, I. J., NIRCHIO, M., OLIVEIRA, C., RON, E. & synonym of Bassobythites, with notes on the synonymy and GAVIRIA, J. 2007. A new species of mullet (Teleostei: distribution of L. brunswigi. Japanese Journal of Ichthyology Mugilidae) from Venezuela, with a discussion of the tax- 38 (2): 125-132. onomy of Mugil gaimardianus. Journal of Fish Biology, 71 COLLETTE, B. B., MCDOWELL, J. R. & GRAVES, J. 2006. (Supplement A): 76-97. Phylogeny of Recent billfishes (Xiphioidei). Bulletin of HAWN, D. R. & COLLETTE, B. B. 2012. 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